Justification:Takifugu niphobles is distributed from the northwest Pacific, from China to Viet Nam, where it is common and can be locally abundant. It is a coastal, shallow water, demersal species which can be found at depths ranging from very shallow waters to 20 m. Takifugu niphobles is an intertidal spawner which is known to form large spawning aggregations on sandy and rubble beaches. There is little population information available, and there are no known species-specific threats. However, the extent to which T. niphobles is impacted by environmental degradation resulting from large-scale anthropogenic activity within its range is largely unknown. There are no known species-specific conservation measures in place for T. niphobles, but it its range overlaps with several marine protected areas. It is therefore listed as Least Concern. Due to the economic importance of the Takifugu genus, and the prevalence of taxonomic uncertainty within this group, we recommend further taxonomic studies utilizing both molecular and morphological methods.

In the Northwest Pacific, Takifugu niphobles is widely distributed in coastal areas from South Korea, Japan, and China to Viet Nam (Duong 2001, Nakabo 2002, Su and Li 2002). It may be found in the Bohai Sea (Su and Li 2002). It is found at depths ranging from 1–20 metres.

Takifugu niphobles is most closely related to T. alboplumbeus (Yamanoue et al. 2008). Due to their relatively recent divergence times, Takifugu interspecific crosses produced by artificial and natural fertilization in both natural and laboratory settings were found to be viable (Fujita 1967, Masuda et al. 1991, Miyaki et al. 1995, Kai et al. 2005). Each combination of Takifugu species is expected to produce fertile hybrid crosses (Yamanoue et al. 2008).Takifugu niphobles is one of the two most common species of puffer fish found in Hong Kong waters, and although traditional fisheries resources have declined drastically in the past decades, T. niphobles can still be easily caught along the seashore (Yu and Yu 2002). Between June 1997 and August 1998, about 100 puffer fish specimens were able to be easily collected from Hong Kong waters every two months for toxicity studies (Yu and Yu 2002). This species is also described as being very common along the coasts of Japan (Honma et al. 1980). Between March 2007 and February 2008, fish sampling was conducted during the spring tide once a month in northwestern Kyushu Island, Japan. Juveniles of this species were a numerically dominant component of the surf zones of sandy beaches, representing 13.6% of the total catch (Inui et al. 2010). It has also been described as as abundant in Cheonsu Bay, Korea (Lee 1995).

Takifugu niphobles is very common in museum collections. It is represented by at least 80 lots (FishNet2 Database searched December 2013).

Takifugu niphobles is most frequently found in shallow areas over sandy or rubble bottoms (Yamahira et al. 1996) at depths ranging from very shallow waters to 20 metres. It is a peripheral freshwater fish, which is often seen in brackish water and is known to occasionally make short trips into small freshwater streams, possibly to rid itself of ectoparasites (Kato et al. 2010). It is a top predator of hard-shelled prey such as molluscs (Shegemiya 2004).

Groups nearing 1,000 individuals have been observed to gather off specific beaches in Japan from late May to early July, annually, to spawn during phases at dusk for several days during spring tides. Males outnumber females during the spawning season, and T. niphobles will form spawning groups consisting of one ripe female and several males. As an intertidal spawner, T. niphobles will deposit eggs and sperm on beaches with rock, pebble, and boulder bottoms. Spawning fish fling themselves out of the water and fertilization occurs on the beach, where gametes are then swept back into the sea, or may be incubated under rocks on the beach, thus enduring periods of exposure to air (Yamahira 1997, K. Matsuura pers. comm. 2011). In Hong Kong waters, the annual spawning season of this species is from October to February (Yu and Yu 2002). The toxicity of the ovaries increases during non-spawning periods, and decreases during spawning.

The ovaries, liver and intestine are extremely toxic. The skin is highly toxic. The flesh and testes are slightly toxic.

The genus Takifugu speciated and radiated in marine waters around China, Korea, and Japan. The highest species density is found in the Bohai Sea, Yellow Sea, and East China Sea, followed by the Sea of Japan and Pacific Coast of Japan, and finally by the South China Sea. Several species have been reported from the Indian Ocean (Yamanoue et al. 2008).

Tetraodontids are characterized by a tough skin that is often covered with small spinulous scales, a beak-like dental plate divided by a median suture, a slit-like gill opening anterior to the base of the pectoral fin, no pelvic fins, no fin spines, a single usually short-based dorsal fin, a single usually short-based anal fin, and no ribs. They are capable of inflating their abdomens with water when frightened or disturbed and are capable of producing and accumulating toxins such as tetrodotoxin and saxitoxin in the skin, gonads, and liver. The degree of toxicity varies by species, and also according to geographic area and season (Allen and Randall 1977, Allen and Erdmann 2012). Fishes in the family Tetraodontidae have the smallest vertebrate genomes known to date (Neafsey and Palumbi 2003).

Takifugu niphobles is incidentally caught but not targeted. In Japan this species is called "Kusa-fugu" (Nakabo 2002). Despite a ban on the sale of toxic pufferfishes in fish markets, T. niphobles may sometimes found in processed fish products in Taiwan (Huang et al. 2014). Takifugu niphobles, like other fugu species,is primarily caught by bottom long-line fishing, a method which is highly effective at landing Takifugu spp. pufferfishes (K. Matsuura pers. comm. 2011).

AquacultureIt is unknown whether T. niphobles is aqua-cultured in parts of its range. Fishes of the genus Takifugu have become the focus of increasing aquaculture efforts throughout East Asia to meet the increasing demand for pufferfish products and to enhance natural populations which have been depleted throughout the region (Kawata et al. 2012).

There have been no confirmed population declines in T. niphobles. However, it is likely to be impacted by the following:

Fugu fishery The Fugu fishery is acknowledged to have undergone significant declines throughout East Asia. Highly effective fishing gear, including modified long-lines and nets with small mesh sizes, rather than excessive fishing effort, have been implicated in the depletion of Takifugu pufferfish resources in parts of East Asia. In Japan, initial efforts to regulate the fishery in the mid-2000s had not achieved desired results by 2010, and were subsequently re-evaluated (Kawata 2012). Although the species-specific effects of the Fugu fishery on T. niphobles are unknown, it is possible that populations of T. niphobles have been affected.

Genetic effects of cultured fish on natural populationsFishes of the genus Takifugu have become the focus of increasing aquaculture efforts throughout East Asia. Aquacultured Takifugu are used to meet increasing demand for pufferfish products and to enhance natural populations which have been depleted throughout the region (Kawata et al. 2012). As culture fish are genetically distinct from natural populations, the release of aqua-cultured fish can result in a range of genetic outcomes, from no detectable effect to complete introgression or displacement of wild populations (Hindar et al. 1991). Fishes of the genus Takifugu are relatively recently diverged, and each combination of Takifugu species is expected to produce fertile hybrid crosses (Yamanoue et al. 2008). It is therefore likely that the effect of intentional and unintentional release of cultured Takifugu on the genetic integrity of wild populations is significant.

Regional threats: environmental degradation and over-fishingMajor threats to biodiversity of the China seas include over-exploitation of fishery resources and environmental deterioration. The China Seas have faced severe environmental degradation due to a range of anthropogenic activities within a relatively recent and short time frame (Daoji and Daler 2004). The degradation of estuarine environments due to pollution and coastal production is of particular concern, as these areas are characterized by high productivity and represent spawning and nursery areas for many species (Liu 2013). Large areas of the China Seas (Liu 2013) and the Gulf of Thailand (Blaber 2000) are considered to be heavily overfished. Additionally, heavy bottom-trawling in the 1980s and the widespread use of modified driftnets for multi-species fisheries in the Bohai Sea, combined with other anthropogenic stresses, have been implicated in the steady decrease in fish landings in this area (Xianshi 2004). In the Yellow Sea, previously dominant large demersal species became the targets of heavy fishing pressure during the 1950s and 1960s and greatly declined in abundance. By the 1980s, many large pelagic species were also showing great declines in abundance, and since that time the dominant species in the Yellow Sea have been small, planktivorous pelagic species, such as anchovies and sardines (Jin and Tang 1996). In the Yellow Sea, all ecological indexes such as the species number, species richness, species diversity and the evenness were lower in the year 2000 than in the year 1985 (Lin et al. 2005).

There are no known species-specific conservation measures in place for T. niphobles, however, it is possible that management efforts aimed at sustaining T. rubripes fisheries have benefited T. niphobles.

Fugu FisheryTakifugu niphobles is not a targeted species in the Fugu fishery, however it is taken incidentally. The Fugu fishery is acknowledged to have undergone significant declines. Japan, South Korea and China previously operated their fisheries under a policy which prevented the regulation of foreign fishing vessels and treated the open sea in East Asia as common property. Since 1999, with the ratification of the United Nations Convention on the Law of the Sea and new fishery agreements between these nations, this policy has shifted to enable coastal countries to regulate foreign fishing vessels. As a result of this policy shift, Japan announced the Plan for Rebuilding Puffer Resources in April 2005, which set dates for an off-fishing season, restrictions on minimum body size, support for stock-enhancement programs, improvements to fishing grounds, and mandated the release of small fishes. As of 2010, stocks had not fully improved, prompting a re-assessment of the program. Several recommendations have been made in order to ensure its continued existence, particularly in light of socio-economic constraints which limit the possibility of developing alternative fisheries. Recommendations include gear restrictions and mandating that fishers catch older and heavier fish by postponing the beginning of the fishing season (Kamara 2012).

In order to sustain fisheries of the East China Sea, the government of China has implemented a number of management and conservation measures. These include establishing a prohibited-fishing zone along the 50-m depth contour, the establishment of seventeen national nature reserves and five special marine protected areas, the creation of fishery protected areas which are annually closed to trawling, and a summer closed-fishing areas, which prohibit trawling and have been extended to the South China sea, Yellow Sea, and Bohai Sea (Cheng et al. 2007).

Further research The economically important genus Takifugu has been recommended for further taxonomic studies based on morphological and molecular analyses (Yamanoue et al. 2008), and we support this recommendation.